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Cellular Respiration Harvesting Chemical Energy. ATP. What’s the point?. The point is to make ATP !. ATP. Harvesting stored energy. Energy is stored in organic molecules carbohydrates, fats, proteins Heterotrophs eat these organic molecules food digest organic molecules to get… - PowerPoint PPT Presentation
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AP Biology
Harvesting stored energy Energy is stored in organic molecules
carbohydrates, fats, proteins Heterotrophs eat these organic molecules food
digest organic molecules to get… raw materials for synthesis fuels for energy
controlled release of energy “burning” fuels in a series of
step-by-step enzyme-controlled reactions
AP Biology
Harvesting stored energy Glucose is the model
catabolism of glucose to produce ATP
C6H12O6 6O2 ATP 6H2O 6CO2+ + +
CO2 + H2O + heatfuel
(carbohydrates)
COMBUSTION = making a lot of heat energy by burning fuels in one step
RESPIRATION = making ATP (& some heat)by burning fuels in many small steps
CO2 + H2O + ATP (+ heat)
ATPglucose
glucose + oxygen energy + water + carbondioxide
res
pir
ati
on
O2 O2
+ heat
enzymesATP
AP Biology
How do we harvest energy from fuels? Digest large molecules into smaller ones
break bonds & move electrons from one molecule to another as electrons move they “carry energy” with them that energy is stored in another bond,
released as heat or harvested to make ATP
e-
+ +e-
+ –loses e- gains e- oxidized reduced
oxidation reduction
redox
e-
AP Biology
How do we move electrons in biology? Moving electrons in living systems
electrons cannot move alone in cells electrons move as part of H atom move H = move electrons
pe
+
H
+H
+ –loses e- gains e- oxidized reduced
oxidation reduction
C6H12O6 6O2 6CO2 6H2O ATP+ + +
oxidation
reductionHe-
AP Biology
Coupling oxidation & reduction REDOX reactions in respiration
release energy as breakdown organic molecules break C-C bonds strip off electrons from C-H bonds by removing H atoms
C6H12O6 CO2 = the fuel has been oxidized electrons attracted to more electronegative atoms
in biology, the most electronegative atom? O2 H2O = oxygen has been reduced
couple REDOX reactions & use the released energy to synthesize ATP
C6H12O6 6O2 6CO2 6H2O ATP+ + +
oxidation
reduction
O
2
AP Biology
Oxidation & reduction Oxidation
adding O removing H loss of electrons releases energy exergonic
Reduction removing O adding H gain of electrons stores energy endergonic
C6H12O6 6O2 6CO2 6H2O ATP+ + +
oxidation
reduction
AP Biology
Moving electrons in respiration Electron carriers move electrons by
shuttling H atoms around NAD+ NADH (reduced) FAD+2 FADH2 (reduced)
+ Hreduction
oxidation
PO–
O–
O
–O
PO–
O–
O
–O
CC
O
NH2
N+
H
adenine
ribose sugar
phosphates
NAD+
nicotinamideVitamin B3niacin
PO–
O–
O
–O
PO–
O–
O
–O
CC
O
NH2
N+
HNADH
carries electrons as a reduced molecule
reducing power!
How efficient!Build once,
use many ways
H
like $$in the bank
AP Biology
Overview of cellular respiration 4 metabolic stages
Anaerobic respiration1. Glycolysis
respiration without O2
in cytosol
Aerobic respiration respiration using O2
in mitochondria
2. Pyruvate oxidation
3. Krebs cycle
4. Electron transport chain
C6H12O6 6O2 ATP 6H2O 6CO2+ + + (+ heat)
AP Biology
ATP synthase enzyme H+ flows through it
conformational changes
bond Pi to ADP to make ATP
set up a H+ gradient allow the H+ to flow
down concentration gradient through ATP synthase
ADP + Pi ATPH+
H+H+
H+
H+ H+
H+H+H+
ATP
ADP P+
But… How is the proton (H+) gradient formed?
And how do we do that?
AP Biology 2006-2007H+
H+H+
H+
H+ H+
H+H+H+
ATP
Got to wait untilthe sequel!
Got the Energy? Ask Questions!
ADP P+
AP Biology
Glycolysis
glucose pyruvate2x6C 3C
In thecytosol?
Why doesthat make
evolutionarysense?
That’s not enoughATP for me!
Breaking down glucose “glyco – lysis” (splitting sugar)
ancient pathway which harvests energy but it’s inefficient
generate only 2 ATP for every 1 glucose occurs in cytosol
AP Biology
Evolutionary perspective Prokaryotes
first cells had no organelles Anaerobic atmosphere
life on Earth first evolved without free oxygen (O2) in atmosphere
energy had to be captured from organic molecules in absence of O2
Prokaryotes that evolved glycolysis are ancestors of all modern life ALL cells still utilize glycolysis
Enzymesof glycolysis are“well-conserved”
AP Biology
10 reactions convert
glucose (6C) to 2 pyruvate (3C)
produces: 4 ATP & 2 NADH
consumes:2 ATP
net yield: 2 ATP & 2 NADH
glucoseC-C-C-C-C-C
fructose-1,6bPP-C-C-C-C-C-C-P
DHAPP-C-C-C
G3PC-C-C-P
pyruvateC-C-C
Overview
DHAP = dihydroxyacetone phosphateG3P = glyceraldehyde-3-phosphate
ATP2
ADP2
ATP4
ADP4
NAD+2
2Pi
enzyme
enzyme
enzyme enzyme
enzyme
enzyme
enzyme
enzyme
2Pi
2H2
AP Biology
Glycolysis summary endergonicinvest some ATP
exergonicharvest a little ATP & a little NADH
net yield2 ATP2 NADH
4 ATP
ENERGY INVESTMENT
ENERGY PAYOFF
G3PC-C-C-P
NET YIELD
like $$in the bank
-2 ATP
AP Biology
Pi
3
6
4,5
ADP
NAD+
Glucose
hexokinase
phosphoglucoseisomerase
phosphofructokinase
Glyceraldehyde 3-phosphate (G3P)
Dihydroxyacetonephosphate
Glucose 6-phosphate
Fructose 6-phosphate
Fructose 1,6-bisphosphate
isomerase
glyceraldehyde3-phosphate
dehydrogenase
aldolase
1,3-Bisphosphoglycerate(BPG)
1,3-Bisphosphoglycerate
(BPG)
1
2
ATP
ADP
ATP
NADH
NAD+
NADH
Pi
CH2
C O
CH2OH
P O
CH2 O P
O
CHOH
C
CH2 O P
O
CHOH
CH2 O PO
CH2OP
O
PO
CH2
H
CH2OHO
CH2 POO
CH2OH
P O
1st half of glycolysis (5 reactions)
Glucose “priming”
get glucose ready to split phosphorylate
glucose molecular
rearrangement split destabilized
glucose
AP Biology
2nd half of glycolysis (5 reactions)
Payola!Finally some
ATP!
7
8
H2O9
10
ADP
ATP
3-Phosphoglycerate(3PG)
3-Phosphoglycerate(3PG)
2-Phosphoglycerate(2PG)
2-Phosphoglycerate(2PG)
Phosphoenolpyruvate(PEP)
Phosphoenolpyruvate(PEP)
Pyruvate Pyruvate
phosphoglyceratekinase
phosphoglycero-mutase
enolase
pyruvate kinase
ADP
ATP
ADP
ATP
ADP
ATP
H2O
CH2OH
CH3
CH2
O-
O
C
PH
CHOH
O-
O-
O-
C
C
C
C
C
C
P
P
O
O
O
O
O
O
CH2
NAD+
NADH
NAD+
NADH
Energy Harvest G3P
C-C-C-PPiPi 6
DHAPP-C-C-C
NADH production G3P donates H oxidizes the sugar reduces NAD+
NAD+ NADH ATP production
G3P pyruvate PEP sugar donates P
“substrate level phosphorylation”
ADP ATP
AP Biology
Substrate-level Phosphorylation
P is transferred from PEP to ADPkinase enzymeADP ATP
I get it!The Pi camedirectly from
the substrate!
H2O9
10
Phosphoenolpyruvate(PEP)
Phosphoenolpyruvate(PEP)
Pyruvate Pyruvate
enolase
pyruvate kinaseADP
ATP
ADP
ATP
H2O
CH3
O-
O
C
O-
C
C
C
P
O
O
O
CH2
In the last steps of glycolysis, where did the P come from to make ATP? the sugar substrate (PEP)
ATP
AP Biology
Energy accounting of glycolysis
Net gain = 2 ATP + 2 NADH some energy investment (-2 ATP) small energy return (4 ATP + 2 NADH)
1 6C sugar 2 3C sugars
2 ATP 2 ADP
4 ADP
glucose pyruvate2x6C 3C
All that work! And that’s all
I get?
ATP4
2 NAD+ 2 Butglucose has
so much moreto give!
AP Biology
Is that all there is? Not a lot of energy…
for 1 billon years+ this is how life on Earth survived no O2 = slow growth, slow reproduction
only harvest 3.5% of energy stored in glucose more carbons to strip off = more energy to harvest
Hard wayto makea living!
O2
O2
O2
O2
O2
glucose pyruvate6C 2x 3C
AP Biology
7
8
H2O9
10
ADP
ATP
3-Phosphoglycerate(3PG)
3-Phosphoglycerate(3PG)
2-Phosphoglycerate(2PG)
2-Phosphoglycerate(2PG)
Phosphoenolpyruvate(PEP)
Phosphoenolpyruvate(PEP)
Pyruvate Pyruvate
ADP
ATP
ADP
ATP
ADP
ATP
H2O
NAD+
NADH
NAD+
NADH
PiPi 6
Glycolysisglucose + 2ADP + 2Pi + 2 NAD+ 2 pyruvate + 2ATP + 2NADH
But can’t stop there!
Going to run out of NAD+
without regenerating NAD+, energy production would stop!
another molecule must accept H from NADH so NAD+ is freed up for another round
PiNAD+
G3P
1,3-BPG 1,3-BPG
NADH
NAD+
NADH
Pi
DHAP
raw materials products
AP Biology
NADH
pyruvate
acetyl-CoA
lactate
ethanol
NAD+
NAD+
NADH
NAD+
NADH
CO2
acetaldehyde
H2O
Krebscycle
O2
lactic acidfermentation
with oxygenaerobic respiration
without oxygenanaerobic respiration
“fermentation”
How is NADH recycled to NAD+?Another molecule must accept H from NADH
recycleNADH
which path you use depends on who you are…
which path you use depends on who you are…
alcoholfermentation
AP Biology
Fermentation (anaerobic) Bacteria, yeast
1C3C 2Cpyruvate ethanol + CO2
Animals, some fungi
pyruvate lactic acid3C 3C
beer, wine, bread
cheese, anaerobic exercise (no O2)
NADH NAD+
NADH NAD+
back to glycolysis
back to glycolysis
AP Biology
recycleNADH
Alcohol Fermentation
1C3C 2Cpyruvate ethanol + CO2
NADH NAD+
Count thecarbons!
Dead end process at ~12% ethanol,
kills yeast can’t reverse the
reaction
bacteria yeast
back to glycolysis
AP Biology
recycleNADH
Reversible process once O2 is available,
lactate is converted back to pyruvate by the liver
Lactic Acid Fermentationpyruvate lactic acid
3C 3CNADH NAD+
Count thecarbons!
O2
animalssome fungi
back to glycolysis
AP Biology
Pyruvate is a branching pointPyruvate
O2O2
mitochondriaKrebs cycle
aerobic respiration
fermentationanaerobicrespiration
AP Biology
H+
H+H+
H+
H+ H+
H+H+H+
And how do we do that?
ATP
But… Have we done that yet?
ADP P+
ATP synthase set up a H+ gradient allow H+ to flow
through ATP synthase powers bonding
of Pi to ADP
ADP + Pi ATP
AP Biology
pyruvate CO2
Glycolysis is only the start Glycolysis
Pyruvate has more energy to yield 3 more C to strip off (to oxidize) if O2 is available, pyruvate enters mitochondria enzymes of Krebs cycle complete the full
oxidation of sugar to CO2
2x6C 3Cglucose pyruvate
3C 1C
AP Biology
intermembranespace inner
membrane
outermembrane
matrix
cristae
Mitochondria — Structure Double membrane energy harvesting organelle
smooth outer membrane highly folded inner membrane
cristae intermembrane space
fluid-filled space between membranes matrix
inner fluid-filled space DNA, ribosomes enzymes
free in matrix & membrane-bound
mitochondrialDNA
What cells would have a lot of mitochondria?
AP Biology
Mitochondria – Function
What does this tell us about the evolution of eukaryotes?
Endosymbiosis!
Dividing mitochondriaWho else divides like that?
Advantage of highly folded inner membrane?More surface area for membrane-bound enzymes & permeases
Membrane-bound proteinsEnzymes & permeases
Oooooh!Form fits function!
bacteria!
AP Biology
pyruvate acetyl CoA + CO2
Oxidation of pyruvate
NAD
3C 2C 1C[2x ] Pyruvate enters mitochondrial matrix
3 step oxidation process releases 2 CO2 (count the carbons!)
reduces 2 NAD 2 NADH (moves e-) produces 2 acetyl CoA
Acetyl CoA enters Krebs cycle
Wheredoes theCO2 go?Exhale!
AP Biology
Pyruvate oxidized to Acetyl CoA
Yield = 2C sugar + NADH + CO2
reduction
oxidation
Coenzyme APyruvate
Acetyl CoA
C-C-CC-CCO2
NAD+
2 x [ ]
AP Biology
Krebs cycle
aka Citric Acid Cycle in mitochondrial matrix 8 step pathway
each catalyzed by specific enzyme step-wise catabolism of 6C citrate molecule
Evolved later than glycolysis does that make evolutionary sense?
bacteria 3.5 billion years ago (glycolysis) free O2 2.7 billion years ago (photosynthesis)
eukaryotes 1.5 billion years ago (aerobic respiration = organelles mitochondria)
1937 | 1953
Hans Krebs1900-1981
AP Biology
4C
6C
4C
4C
4C
2C
6C
5C
4C
CO2
CO2
citrate
acetyl CoACount the carbons!
3Cpyruvate
x2
oxidationof sugars
This happens twice for each glucose molecule
AP Biology
4C
6C
4C
4C
4C
2C
6C
5C
4C
CO2
CO2
citrate
acetyl CoACount the electron carriers!
3Cpyruvate
reductionof electron
carriers
This happens twice for each glucose molecule x2
CO2
NADH
NADH
NADH
NADH
FADH2
ATP
AP Biology
So we fully oxidized glucose
C6H12O6
CO2
& ended up with 4 ATP!
Whassup?
What’s the point?
AP Biology
Krebs cycle produces large quantities of electron carriers NADH FADH2
go to Electron Transport Chain!
Electron Carriers = Hydrogen Carriers
What’s so important about
electron carriers?
H+
H+H+
H+
H+ H+
H+H+H+
ATP
ADP+ Pi
AP Biology
Energy accounting of Krebs cycle
Net gain = 2 ATP
= 8 NADH + 2 FADH2
1 ADP 1 ATPATP
2x
4 NAD + 1 FAD 4 NADH + 1 FADH2
pyruvate CO2
3C 3x 1C
AP Biology
Value of Krebs cycle? If the yield is only 2 ATP then how was the
Krebs cycle an adaptation? value of NADH & FADH2
electron carriers & H carriers reduced molecules move electrons reduced molecules move H+ ions
to be used in the Electron Transport Chain
like $$in the bank
AP Biology
H+
H+H+
H+
H+ H+
H+H+H+
And how do we do that?
ATP
But… Have we done that yet?
ADP P+
ATP synthase set up a H+ gradient allow H+ to flow
through ATP synthase powers bonding
of Pi to ADP
ADP + Pi ATP
AP Biology
ATP accounting so far… Glycolysis 2 ATP Kreb’s cycle 2 ATP Life takes a lot of energy to run, need to
extract more energy than 4 ATP!
A working muscle recycles over 10 million ATPs per second
There’s got to be a better way!
I need a lotmore ATP!
AP Biology
There is a better way! Electron Transport Chain
series of proteins built into inner mitochondrial membrane along cristae transport proteins & enzymes
transport of electrons down ETC linked to pumping of H+ to create H+ gradient
yields ~36 ATP from 1 glucose! only in presence of O2 (aerobic respiration)
O2That
sounds morelike it!
AP Biology
Mitochondria Double membrane
outer membrane inner membrane
highly folded cristae enzymes & transport
proteins intermembrane space
fluid-filled space between membranes
Oooooh!Form fits function!
AP Biology
Electron Transport Chain
Intermembrane space
Mitochondrial matrix
Q
C
NADH dehydrogenase
cytochromebc complex
cytochrome coxidase complex
Innermitochondrialmembrane
AP Biology
G3PGlycolysis Krebs cycle
8 NADH2 FADH2
Remember the Electron Carriers?
2 NADH
Time tobreak open
the piggybank!
glucose
AP Biology
Electron Transport Chain
intermembranespace
mitochondrialmatrix
innermitochondrialmembrane
NAD+
Q
C
NADH H2O
H+
e–
2H+ + O2
H+H+
e–
FADH2
12
NADH dehydrogenase
cytochromebc complex
cytochrome coxidase complex
FAD
e–
H
H e- + H+
NADH NAD+ + H
H
pe
Building proton gradient!
What powers the proton (H+) pumps?…
AP Biology H+
H+H+
H+
H+ H+
H+H+H+
ATP
NAD+
Q
C
NADH H2O
H+
e–
2H+ + O2
H+H+
e–FADH2
12
NADH dehydrogenase
cytochrome bc complex
cytochrome coxidase complex
FAD
e–
Stripping H from Electron Carriers Electron carriers pass electrons & H+ to ETC
H cleaved off NADH & FADH2
electrons stripped from H atoms H+ (protons) electrons passed from one electron carrier to next in
mitochondrial membrane (ETC) flowing electrons = energy to do work
transport proteins in membrane pump H+ (protons) across inner membrane to intermembrane space
ADP+ Pi
TA-DA!!Moving electrons
do the work!
H+ H+ H+
AP Biology
But what “pulls” the electrons down the ETC?
electronsflow downhill
to O2 oxidative phosphorylation
O2
H2O
AP Biology
Electrons flow downhill Electrons move in steps from
carrier to carrier downhill to oxygen each carrier more electronegative controlled oxidation controlled release of energy
make ATPinstead of
fire!
AP BiologyH+
ADP + Pi
H+H+
H+
H+ H+
H+H+H+We did it!
ATP
Set up a H+
gradient Allow the protons
to flow through ATP synthase
Synthesizes ATP
ADP + Pi ATP
Are wethere yet?
“proton-motive” force
AP Biology
The diffusion of ions across a membrane build up of proton gradient just so H+ could flow
through ATP synthase enzyme to build ATP
Chemiosmosis
Chemiosmosis links the Electron Transport Chain to ATP synthesis
Chemiosmosis links the Electron Transport Chain to ATP synthesis
So that’sthe point!
AP Biology
Peter Mitchell Proposed chemiosmotic hypothesis
revolutionary idea at the time
1961 | 1978
1920-1992
proton motive force
AP Biology
H+
H+
O2+
Q C
ATP
Pyruvate fromcytoplasm
Electrontransportsystem
ATPsynthase
H2O
CO2
Krebscycle
Intermembranespace
Innermitochondrialmembrane
1. Electrons are harvested and carried to the transport system.
2. Electrons provide energy
to pump protons across the membrane.
3. Oxygen joins with protons to form water.
2H+
NADH
NADH
Acetyl-CoA
FADH2
ATP
4. Protons diffuse back indown their concentrationgradient, driving the synthesis of ATP.
Mitochondrial matrix
21
H+
H+
O2
H+
e-
e-
e-
e-
ATP
AP Biology
Summary of cellular respiration
Where did the glucose come from? Where did the O2 come from? Where did the CO2 come from? Where did the CO2 go? Where did the H2O come from? Where did the ATP come from? What else is produced that is not listed
in this equation? Why do we breathe?
C6H12O6 6O2 6CO2 6H2O ~40 ATP+ + +
AP Biology
ETC backs up nothing to pull electrons down chain NADH & FADH2 can’t unload H
ATP production ceases cells run out of energy and you die!
Taking it beyond… What is the final electron acceptor in
Electron Transport Chain?
O2
So what happens if O2 unavailable?
NAD+
Q
C
NADH H2O
H+
e–
2H+ + O2
H+H+
e–FADH2
12
NADH dehydrogenase
cytochrome bc complex
cytochrome coxidase complex
FAD
e–